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Sperm competition and male forceps dimorphism in the European earwig Forficula auricularia (Dermaptera: Forficulina) Gordon S. Brown A thesis submitted to the University of St. Andrews in application for the degree of Doctor of Philosophy Submitted: 6 October 2006 Supervisor: Dr. Joseph L. Tomkins Co-supervisor: Prof. Michael G. Ritchie Declaration I, Gordon S. Brown, hereby certify that this thesis, which is approximately 25000 words in length, has been written by me, that it is the record of work carried out by me and that it has not been submitted in any previous application for a higher degree. Date ……………… signature of candidate ……………………………… I was admitted as a research student in August 2002 and as a candidate for the degree of PhD in September 2003; the higher study for which this is a record was carried out in the University of St Andrews between 2002 and 2006. Date ……………… signature of candidate ……………………………… In submitting this thesis to the University of St Andrews I understand that I am giving permission for it to be made available for use in accordance with the regulations of the University Library for the time being in force, subject to any copyright vested in the work not being affected thereby. I also understand that the title and abstract will be published, and that a copy of the work may be made and supplied to any bona fide library or research worker. Date ……………… signature of candidate ……………………………… I hereby certify that the candidate has fulfilled the conditions of the Resolution and Regulations appropriate for the degree of PhD in the University of St Andrews and that the candidate is qualified to submit this thesis in application for that degree. Date ……………… signature of supervisor …………………………..… Contents Acknowledgements Abstract Chapter 1: Introduction Chapter 2: Geographic variation of the male dimorphism in UK populations of the European earwig Chapter 3: Development of multi-purpose molecular genetic markers for the European earwig Chapter 4: Phylogeography of the European Earwig in the UK Chapter 5: Ejaculate characteristics of European earwig Chapter 6: Concluding remarks Appendices A. Specimens collected B. Population density estimates through trapping C. Published work D. DNA extraction E. Processing raw microsatellite data from a Beckman Coulter DNA sequencer F. Summary of genetic dataset References 3 The hornie gollach is an awesome beast soople and scaly, it has twa horns and a hantle o feet and a forkie tailie (Burnett 1937) 4 Acknowledgements I have many people to thank for helping me through the past four years. Firstly, I thank my family without whose love and support I doubt I would have made it as far as doing a PhD. To Joe, my long-suffering supervisor, my thanks for introducing me to the wonderful world of the earwig and for helping me through even when time and distance made it difficult. I would also like to thank Mike Ritchie standing in when Joe's destiny took him to the warmer climes of Perth, for his insights into my murky population genetic analysis and, indeed, for putting me in touch with Joe in the first place. My thanks also to Jeff Graves for sharing his experience in the lab and in the world of genetics whilst spreading his omnipresent unflappable calm in the face of truculent molecules. Many thanks to Tash Lebas for sharing her knowledge and advice, particularly with regard to taming those aforesaid truculent molecules. My time in St. Andrews would have been a bland and mirthless experience if hadn't been for the excellent friends I have made. In particular I will always remember with a smile the pub-quiz and sublimely ridiculous nights at home with Nathan, Paty, Kirsten, Susi and Oli. I have also enjoyed a good laugh with the myriad friendly bodies about the “Harold Mitchell”. In particular thanks to my friends; Valentina, Joe Milton, Vicki, Emma, Mark Chapman, Mark Lineham, Veronica, Kathryn and Marcos with whom I've been fortunate to share an office. I would also like to thank the many other people who helped make the extended community of the university so welcoming and enjoyable to be a part of; Peter Slater whose warm welcome to St. Andrews at my interview I will always remember, to Lucy & her 'darling' my thanks for good times and the generous gift of a washing machine, to Ian Matthews, Derwent, Betsy, Lorraine, Jared, Nicky, Julian, Mark Hulme and Miriam and all the rest. To my friends Sally and Robin and their fine vessel, Tub; thank you for helping me to escape St Andrew to the finer parts of Scotland; something I aim to do a lot more of in the future and I hope we will have many more opportunities to sail together in the future. 5 To friends from a past life; Peter and Liz Gibson, Graham, Alex, Antonis, Rachel, Tony and Martin and other friends in Edinburgh. The good times I spent there have been influential in finding my path to where I am now. Hopefully our paths will cross again before too long. To the folk who help oil the machinery of the labs in the HMB; Tanya for maintaining the lab environment as an enjoyable place to work and to Harry Hodge and Ray Stephenson for helping me with my occasional requests for random bits of lab equipment. From my challenging time associated with “The Beckman” I would like to offer my thanks to; Tom Meagher for granting access to his machine, to Veronica Poland for her introduction and shared woes, to Bill Hutchinson who shared his approach to processing the Beckman data which lead to the improvement of my own method. Thanks also to technician Dave Forbes for his efforts maintaining the fickle machinery. I am grateful for the jovial lab environment created and maintained by Joe’s band of merry lab workers and past students; Joolz , Clare, Andy, Lukasz and Piotr. Also, thanks to Cate Lyon for help with DNA extractions. For lab advice from afar I would also like to thank; Matt Gage for sharing his experience of sperm under the microscope and recommending the recipe for Barth’s solution; Jason Kennington (UWA) for advice on PCR and genetic analysis channelled through Joe and Marie Hale for help with the microsats and primer note. I would like that thank and man I have never met, Jeff Glaubitz, for the very useful documentation of his data file utility CONVERT (Glaubitz 2004). This goes far beyond the remit of Glaubitz’s own program and, indeed, beyond the meagre documentation available for a program MICROSAT that I relied upon (Minch 1997). During my PhD I spent many days visiting islands and touring around the British countryside hunting for the (sometimes elusive and at others multitudinous) earwig. I am grateful for having had the opportunity to get to these remote and beautiful parts of the UK and to the people that made this possible. Joe and I made many a trip to the Farnes where Steely, Alien and the other wardens always made it a rewarding experience. I'd like to thank John Walton and the National Trust for granting access to these islands. 6 I addition to the Farne islands I also offer my thanks to the many other landowners and organisations who granted me access to some very special places: Julie Love and Dave Mawer of the Isles of Scilly Wildlife trust; to Mr Dorrien-Smith for access to his estate on Tresco (Isles of Scilly), the Landmark trust for access to Lundy, to Ian Bullock, Simon Avery, Juan Brown and Graham Thompson of the Countryside commission for Wales for access to the Pembrokeshire islands of Ramsey and Skomer and samples from Skokholm. In the Firth of Forth I visited the islands of Inchcolm, Inchkeith, Fidra and Inchmickery, Lamb and Eyebroughy, Inch Garvie and Craigleith, with thanks to Ron Selley of Historic Scotland, Paul Barry, Kirsty Smith (RSPB), Baron Camilo Agasim-Pereira of Fulwood, Sir Jack Steward-Clark and Sir Hew Hamilton Dalrymple, respectively. I would also like to thank the individuals who personally helped me during my field work, in particular the IoSWT volunteers, Connie and Pip and Richard Farr and his brother-in-law for a very civilised days work on St. Agnes, Annet and Gugh (the pub lunch is a happy lingering memory). My thanks go out to a small group of people who, in taking the time to go out, collect earwigs and carefully send them to me in St. Andrews, without whose efforts my data and my thesis would be significantly less than it is. These fine people are Joe's parents, Stephen & Helga Tomkins, Tom and Nils Tregenza and Noam Hosken. Finally, I would like to thank NERC for the studentship I received from them and my examiners Chris Todd and Matt Gage for giving up their time to examine my thesis. 7 Abstract The European earwig exhibits a remarkable male-dimorphism in forceps morphology that is associated with alternative reproductive tactics under the control of a conditional evolutionarily stable strategy. Populations on the small, rocky islands of the Farnes off the Northumberland coast are known to sustain populations with dramatically higher morph ratios than observed on the UK mainland. A survey conducted of island and mainland sites around the UK showed that the dimorphic populations of the Farnes are similar to other islands and that mainland populations generally exhibit low morph ratios. Additionally, a correlation between morph ratio and population density was found lending support to the hypothesis that the ESS thresholds that define the morph ratios have diverged through local adaptation. A set of seven microsatellite markers are presented that were developed from a Farne island population of F.
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